Fungal diversity on fallen leaves of Ficus in northern Thailand§

Fungal diversity on fallen leaves of Ficus in northern Thailand § § 1 first-author 2 corresp ‡Corresponding Author ‡Corresponding Author †E-mail:kdhyde1@gmail.comfuchenglin@zju.edu.cn †E-mail:kdhyde1@gmail.comfuchenglin@zju.edu.cn kdhyde1@gmail.com 3 1 corresp ‡Corresponding Author ‡Corresponding Author †E-mail:kdhyde1@gmail.comfuchenglin@zju.edu.cn †E-mail:kdhyde1@gmail.comfuchenglin@zju.edu.cn kdhyde1@gmail.com Biotechnology Institute, Zhejiang University, Hangzhou 310029, China Biotechnology Institute, Zhejiang University, Hangzhou 310029, China Fungal Research Group, School of Science, Mae Fah Luang University, Tasud, Chiang Rai 571000, Thailand Fungal Research Group, School of Science, Mae Fah Luang University, Tasud, Chiang Rai 571000, Thailand Faculty of Agricultural Technology, King Mongkut’s Institute of Technology Ladkrabang, Ladkrabang, Bangkok 10520, Thailand Faculty of Agricultural Technology, King Mongkut’s Institute of Technology Ladkrabang, Ladkrabang, Bangkok 10520, Thailand Fallen leaves of Ficus altissima , F . virens , F . benjamina , F . fistulosa and F . semicordata , were collected in Chiang Mai Province in northern Thailand and examined for fungi. Eighty taxa were identified, comprising 56 anamorphic taxa, 23 ascomycetes and 1 basidiomycete. Common fungal species occurring on five host species with high frequency of occurrence were Beltraniella nilgirica , Lasiodiplodia theobromae , Ophioceras leptosporum , Periconia byssoides and Septonema harknessi . Colletotrichum and Stachybotrys were also common genera. The leaves of different Ficus species supported diverse fungal taxa, and the fungal assemblages on the different hosts showed varying overlap. The fungal diversity of saprobes at the host species level is discussed. Ficus Fallen leaves Saprobes Fungal diversity INTRODUCTION 1 Fungi are chemoheterotrophic organisms that play an important role as decomposers in natural ecosystems (Duarte et al., 2006 2006 ; Gadd, 2007 2007 ). Fungi have additional economic importance as biocontrol agents, chemical producers of bioactive compounds used in the pharmaceutical and many other industries (Yuen et al., 1999 1999 ; Bucher et al., 2004 2004 ; Duarte et al., 2006 2006 ). The economic impact of fungi in the environment is likely related to the total fungal species pool (Hyde, 2001 2001 ; Piepenbring, 2007 2007 ). Research on fungal diversity therefore provides a basis for estimating the functional role of fungi in ecosystems, exploiting these economic fungi and estimating the total fungal species richness in a region or globally (Arnold et al., 2001 2001 ; Mueller and Schmit, 2007 2007 ). Although the 1.5 million species estimated by Hawksworth ( 2001 2001 ) are commonly accepted (Hyde, 2001 2001 ; Hyde et al., 2007 2007 ), the actual quantity of fungal species is unclear (Arnold et al., 2001 2001 ; Hyde, 2001 2001 ; Hyde et al., 2007 2007 ; Schmit and Mueller, 2007 2007 ). Schmit and Mueller ( 2007 2007 ) estimated that there is a minimum of 712 000 fungal species worldwide. This estimate was primarily based on the observed ratio between vascular plant species diversity and fungal diversity in a certain area. Meta-analysis of fungal diversity in small plots (Schmit et al., 2005 2005 ) demonstrated that the numbers of tree species and macrofungal species diversities are correlated (Mueller et al., 2007 2007 ). The actual number of fungi is still unknown; however, only 5%~13% of the total estimated global fungal species have been described. The undescribed fungi may occur in poorly studied countries, hosts, habitats, niches, or tissues, and are mostly microfungi (Hyde, 2001 2001 ; Schmit and Mueller, 2007 2007 ). To determine the accuracy of fungal estimates, more information is needed concerning species diversity in poorly studied areas and hosts, especially in poorly studied areas (Dulymamode et al., 2001 2001 ; Hyde, 2001 2001 ). There have been several studies on fungi in poorly studied habitats in Thailand (Jones and Hyde, 2004 2004 ). This has included studies on larger fungi in the forests of northern Thailand (Le et al., 2007a 2007a ; 2007b 2007b ), fungi on peat swamp palms (Pinnoi et al., 2006 2006 ; Pinruan et al., 2007 2007 ), fungi on other monocotyledons (Photita et al., 2001 2001 ; Thongkantha et al., 2008 2008 ), fungi on leaf litter of angiosperms (Promputtha et al., 2004 2004 ; 2005 2005 ; Duong et al., 2006 2006 ; 2008 2008 ), and fungi on decaying wood (Kodsueb et al., 2008a 2008a ; 2008b 2008b ). There has been, however, no study on the litter of Ficus species. Whether fungi are host-specific or generalists are significant in estimating species numbers (Zhou and Hyde, 2001 2001 ; Hyde et al., 2007 2007 ), is particularly important in studying leaf litter. In tropical forests, plant litter comprises a mixture of diverse host leaves, unlike that in many temperate forests where stands contain low tree diversity. Several studies on the saprobic microfungal diversity associated with plants have been carried out (Hyde et al., 2001 2001 ; Yanna et al., 2001 2001 ; Dulymamode et al., 2001 2001 ; Paulus et al., 2006a 2006a ; 2006b 2006b ), and to a large extent, fungi tend to be host-specific at the host genus level (Taylor et al., 2001 2001 ). Paulus et al.( 2006a 2006a ; 2006b 2006b ) studied 6 tree hosts from 4 families and found very little overlap. The low overlap may be because the hosts studied were from different families. The next question to ask is whether litter of different species of the same host genus has host-specific fungi or generalists? We, therefore, chose to study the saprobes on 5 species of Ficus to establish data on the fungal communities involved in the decay of litter from different species of the same host genus. The genus Ficus (Moraceae) includes some 750 species of woody plants occurring in most tropical and subtropical forests throughout the world (Weiblen, 2000 2000 ). Ficus is an evergreen or deciduous tree, frequently growing on other trees (strangling) with smooth pale gray bark and abundant white latex (Gardner et al., 2000 2000 ). There is little known concerning the fungal diversity on this host genus (Suryanarayanan and Vijaykrishna, 2001 2001 ; Paulus et al., 2006a 2006a ). Paulus et al.( 2006a 2006a ; 2006b 2006b ) reported on the fungal diversity in leaf litter of 2 species of Ficus in Australia, while Suryanarayanan and Vijaykrishna ( 2001 2001 ) isolated 28 endophyte fungi from Ficus benghalensis . Ficus is the second largest tree genus in northern Thailand after Syzygium with 30 species of Ficus growing in the region (Gardner et al., 2000 2000 ). The purpose of the present study is to assess the diversity of saprobic fungi on 5 species of Ficus in northern Thailand in order to (1) investigate fungal distribution on saprobic fallen leaves of Ficus species; and (2) evaluate the different fungal communities involved in the decay of litter from different species of the same host genus. MATERIALS AND METHODS 1 Survey design 2 To examine the effect of host species on microfungal assemblages, 5 species of Ficus were chosen for the study. Individual trees of each species were located at each of the sites selected for sampling (Table 1 1 ). Thirty fallen leaves at various stages of decay were haphazardly collected from each Ficus species from each site within 2 d. All samples were placed in Ziplock plastic bags containing tissue paper moistened with sterile distilled water and sealed and incubated at room temperature. Material was examined in the laboratory on Days 5, 15 and 25 after sampling. Plant sepcies Location Ficus altissima MRC, T. Pa Pae, A. Mae Taeng, Chiang Mai Province, Thailand, 19°07′12″ N, 98°44′2.64″ E. F . virens Out door of MRC, T. Pa Pae, A. Mae Taeng, Chiang Mai Province, Thailand, 19°07′12″ N, 98°44′2.64″ E. F . benjamina About 5 km east of MRC, M. Mae Taeng, Chiang Mai Province, Thailand, 19°05′12″ N, 98°44′2.64″ E. F . fistulosa Near Mae Malai, M. Mae Taeng, Chiang Mai Province, Thailand, 19°05′0″ N, 98°5′42″ E. F . semicordata Tung Cho National Park, forest trail, Tung Joaw Village, Mae Teng Dist., Chiang Mai Province, Thailand, 19°08′4.2″ N, 98°38′ 54″ E. Assessment of microfungal diversity 2 For the assessment of fungal presence, all leaves were cut into 5 cm×5 cm quadrates for observation. Fruiting bodies of microfungi were observed under a stereoscope. A slide was then prepared for one representative fruiting structure of each taxon and placed in water and observed under a compound microscope. Micrographs were taken and stains were added as appropriate. Slides were made semipermanent by the addition of 90% (v/v) lactic acid. Species identifications were made using morphological characters. Herbarium specimens and slides of all taxa are maintained at the Mushroom Research Foundation Herbarium with some duplicates in Zhejiang University Herbarium. Definitions and statistical analyses 2 Taxa were recorded as either present or absent from each leaf. The number of leaves, on which a particular fungal species was observed, designated the ‘occurrence of a fungus’ and was used to calculate the ‘percentage occurrence’ of a taxon on leaves of one tree species using the following formula (Tsui et al., 2001 2001 ; Yanna et al., 2002 2002 ): Percent occurrence of taxon A (%)=occurrence of taxon A/occurrence of all taxa in one tree species×100. Fungal species diversity on each Ficus species was calculated using Shannon-Wiener’s index H : H =−∑ P i ln P i , where P i is the frequency of fungal species i occurring on specific host leaves (Begon et al., 1993 1993 ; Wong and Hyde, 2001 2001 ). Sørensen’s index of similarity ( S ) was applied (Magurran, 1988 1988 ; Tsui et al., 2001 2001 ) to compare the similarity of the species on leaves of different Ficus spp.: S =2 c /( a + b ), where a is the total number of species on host A leaves, b is the total number of species on host B leaves, and c is the number of species on both host leaves. Similarity is expressed with values between 0 (no similarity) and 1 (absolute similarity). RESULTS 1 Species richness and dominant fungi on Ficus spp. 2 Examination of decaying leaves of 5 species of Ficus from northern Thailand yielded 80 fungal taxa, comprising 56 anamorphic taxa, 23 ascomycetes and 1 basidiomycete. We have identified many taxa to species level, although in several cases the morphology of the taxa in our study was not identified to species (unidentified species, Table 2 2 ). Five species, Beltraniella nilgirica , Lasiodiplodia theobromae , Ophioceras leptosporum , Periconia byssoides and Septonema harknessi were common species occurring on all five host species ( > 3% occurrence, Table 2 2 ). In addition, Colletotrichum and Stachybotrys were common genera, with several species occurring on the Ficus leaves (Table 2 2 ). Figs. 1 1 ~ 5 5 also show some rare fungal taxa on the different host species. FV: Ficus virens ; FF: F . fistulosa ; FA: F . altissima ; FS: F . semicordata ; FB: F . benjamina Fungus name FV FF FA FS FB Ascomycetes   Amphisphaeria fallax 2.3   Anthostomella limitata 2.2 2.4   Anthostomelle nannorrhopis 2.3 3.4 5.8   Chaetomium sulphureum 4.7   Diaporthe adunca 4.7   Diaporthe perjuncta 2.4 3.4   Eutypella scoparia 2.4 3.4   Glomerella acutata   Glomerella cingulata 1.1 1.2   Guignardia spp. 1.1 2.4 1.2 2.9   Hyponectria populi 3.4   Kirschsteiniothelia spp. 1.7   Lasiosphaeria breviseta 4.4   Lophiostoma minosporum 6.8   Massarina hepaticarum 4.7   Melanospora brevirostris 3.4   Munkovalsaria appendenta 1.1 1.7 1.4   Mycoenterolobium platysporium 2.2 2.3   Mycosphaerella dianthi 4.8 2.9   Mycosphaerella freycineitae 4.4 3.4   Nectria sinopica 1.2   Ophioceras leptosporum 4.4 4.8 4.7 6.8 5.8   Rhytisma acerinum 3.4 Mitosporic fungi   Acremonium charticola 2.3   Ajrekarella polychaetriae 2.2   Alternaria alternata 4.8 4.3   Beltraina querna 1.2   Beltraniella nilgirica 4.4 4.8 3.6 6.8 5.8   Cercospora apii 2.2 2.9   Chaetospermum artocarpi 4.8   Chaetospermum chaetosporum 5.8   Cladosporium cladosporioides 3.3 3.6 6.8   Colletotrichum dematium 2.2 4.8 3.6 2.9   Colletotrichum gloeosporioides 3.6   Colletotrichum graminicola 4.7   Cylindrocolla urticae 5.8   Dicyma state of Ascotricha chartarum 2.2 2.3   Dinemasporium strigosum 2.2 4.8 2.9   Discosia artocreas 2.4   Discosia brasiliensis 1.4   Ellisiopsis gallesiae 2.3   Fusariella hughesii 4.4 4.7   Fusariella obstipa 2.3   Fusarium chlamydosporum 4.8   Fusarium oxysporum 5.5   Fusarium solani 4.4   Giulia tenuis 1.7   Kellermania yuccifolia 2.4   Kontospora halophila 1.7   Koorchaloma jamaicensis 2.4   Lasiodiplodia theobromae 3.3 4.8 3.6 6.8 5.8   Menispora ciliata 3.4   Menispora glauca 3.4   Microdochium phragmitis 4.8 4.7   Mycoenterolobium platysporum 2.2 2.3   Periconia byssoides 4.4 4.8 4.7 6.8 4.3   Pestalotiopsis gigas 2.2 2.9   Pestalotiopsis owenii 4.4 2.9   Phaeoisaria clematidis 2.3   Phoma eupyrena 5.8   Phoma valerianae 2.2 2.4   Phomopsis asparagi 2.2 1.2 1.4   Plectophomella nypae 2.2   Sagenomella striatispora 2.4   Schwarzmannia goebeliae 1.1   Septonema harknessi 4.4 4.8 4.7 5.1 5.8   Sphaeridium candidum 2.4 2.9   Stachybotrys bisbyi 2.9   Stachybotrys nephrospora 2.9   Stachybotrys oenanthes 4.4 2.3 2.9   Stachybotrys renispora 3.3   Stachybotrys sansevieriae 2.3 3.4   Torula graminis 5.8   Torula herbarum 4.4   Trichothecium roseum 2.4 3.6 3.4   Volutella ciliata 4.4 3.6 6.8   Wiesneriomyces javanicus 2.2 4.8 3.4   Xylohypha nigrescens 4.7   Zygosporium chartarum 2.9 Basidiomycete   Halocyphina spp. 1.2 Fungal diversity 2 Communities of fungal taxa on different Ficus species leaf litter were relatively distinct. The number of taxa varied from 24 ( F . semicordata ) to 33 ( F . virens and F . altissima ) (Table 3 3 ). The fungal communities on F . virens and F . altissima were the most similar but were different from those on F . semicordata (Table 4 4 ). The percentages of ascomycetes on F . virens and F . altissima were similar (24% and 30%, respectively). F . semicordata had the highest percentage of ascomycetes (46%) among the host species. The percentages of anamorphic taxa on F . virens and F . altissima were also similar. 2 Ficus species 3 Number of species 2 Total 2 Index H Ascomycetes Mitosporic fungi Basidomycetes Ficus virens 8 25 33 3.3525 F . fistulosa 6 19 25 3.0887 F . altissima 10 22 1 33 3.3196 F . semicordata 11 13 24 3.0881 F . benjamina 5 21 26 3.1496 Ficus virens F . fistulosa F . altissima F . semicordata F . benjamina Ficus virens 0.41 0.45 0.35 0.44 F . fistulosa 0.31 0.41 0.43 F . altissima 0.35 0.31 F . semicordata 0.28 F . benjamina DISCUSSION 1 Studies of fungal diversity on Ficus have not been carried out until recently, although this genus is very common in forests throughout the world (Weiblen, 2000 2000 ; Paulus et al., 2006a 2006a ; 2006b 2006b ). Suryanarayanan and Vijaykrishna ( 2001 2001 ) studied fungal endophyte diversity on different host tissues of F. benghalensis in India. They found little overlap between fungal endophyte composition on the leaves and aerial roots. Paulus et al.( 2006a 2006a ; 2006b 2006b ) studied saprobic microfungi in leaf litter of Australia tropical rainforests, including two Ficus species. They found that fungal assemblages of F. destruens and F. pleurocarpa grouped closely together. The overlapping species were Beltraniella portoricensis, Beltrania cf. concurvispora, Cylindrocladium coulhounii, Dictypchaeta cf. novae-guineensis, Cylindrosympodium cryptocaryae, Idriella acerosa, Idriella lunata, Menisporopsis theobromae, Ophiognomonia elasticae, Parasypodiella elongate and Trichoderma viride. In the present study, we investigated fungal diversity on leaves of five Ficus species. Our results show that the overlap of fungal species was much higher than that in previous studies on fallen leaves from different hosts in different families. Similarity ranged from 28% to 45%. Some species, such as Ophioceras leptosporum, Beltraniella nilgirica, Lasiodiplodia theobromae, Periconia byssoides and Septonema harknessi, were shared by all hosts with high occurrence, indicating that a ‘core fungal group’ might be responsible for the decay of Ficus leaves (Wong and Hyde, 2001 2001 ). In this study some fungi are new records on Ficus. Among these, many species are widespread, such as Diaporthe adunca, Lasiosphaeria breviseta, Colletotrichum gloeosporioides and Torula herbarum. At least one species, Munkovalsaria appendenta, is a new record for Thailand. Further studies are needed to find more fungi on the leaves of Ficus, for our investigation was limited, with one tree of each host being sampled once. Some previous research results showed that the hosts or tissues are important factors that influence the fungal communities (Ho et al., 2001 2001 ; Paulus et al., 2003 2003 ; Rambelli et al., 2004 2004 ) and that some saprobes might be host-specific (Hyde et al., 1997 1997 ). Wong and Hyde ( 2001 2001 ) investigated fungal diversity on Gramineae , and their results showed that fungal composition on host species level varied. Studies on microfungi of tree leaves in tropical regions suggested that the fungal assemblages on different host species were variable, and some other terms, such as ‘host exclusivity’, ‘host fidelity’, ‘host affinity’, ‘host affinity’, ‘host preference’ and ‘host recurrence’, were used to describe plant-fungal associations with definitive host (Zhou and Hyde, 2001 2001 ). Taylor et al.( 2001 2001 ) noted that saprobic fungi were specific at a host genus level. 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Host-specificity, host-exclusivity and host-recurrence in saprobic fungi We are grateful to Dr. Dhanasekaran Vijaykrishna, University of Hong Kong, China and Dr. Ruilin Zhao, Southwest Forestry University, China for technical help. 10.1631/jzus.B0860005 Project supported by the Mushroom Research Centre, Chiang Mai, Thailand and the National Natural Science Foundation of China (No. 30670072) and the Natural Science Foundation of Zhejiang Province of China (No. Y350568) 2008-07-28 2008-08-15 2022-01-05 Copyright © 2008, Journal of Zhejiang University Science 2008 Journal of Zhejiang University-SCIENCE B (Biomedicine and Biotechnology) 10 09